WAT E R F O W L M A N AG E M E N T H A N D B O O K
13.2.7. Identifying the
Factors That Limit
Duck Production
James K. Ringelman
Colorado Division of Wildlife
317 West Prospect Road
Fort Collins, CO 80526
Low duck populations in the late 1980’s and
early 1990’s prompted unprecedented action from
the natural resources community. Agencies and
private organizations that were traditionally
involved with waterfowl management redoubled
their efforts, in the process forming partnerships
with groups that were relatively new to the
waterfowl management arena. Many resource
managers who have had relatively little experience
with waterfowl habitat management now find
themselves expected to manage duck populations
for increased production. Decades of waterfowl
research and management experience have
provided them with many potential management
tools. Unfortunately, the absence of general
guidelines for directing waterfowl management
actions has put these newcomers to the field at a
decided disadvantage. This is particularly true for
managers who reside outside of the northern Great
Plains, a region that has been the focus of most
research on breeding ducks.
This leaflet is intended to orient managers to
approaches for identifying the factors that limit
duck production. The concepts presented here will
assist in making logical management choices in
regions where little is known about breeding ducks
and their habitat. Although it may serve as interim
guidance, this leaflet is not intended to substitute
Fish and Wildlife Leaflet 13.2.7. • 1992
for rigorous, scientific research on waterfowl
biology. Readers are urged to use this leaflet as a
starting point from which to gather additional
knowledge using companion leaflets and technical
publications.
The Reproductive Cycle
Although ducks are a diverse group of birds,
many dabbling and diving ducks in North America
show similarities in general facets of their breeding
biology. A basic understanding of the important
events and forces that drive reproductive behavior
is essential to interpreting premanagement
information. The following sections provide a
summary of duck breeding biology that, although
not strictly accurate for any particular species, is
generally representative of the most common
North American ducks.
Resource Needs
Most ducks arrive on their breeding grounds
from late March to early May. Shortly thereafter
they begin to make regular use of wetlands that
vary in size, water permanency, and vegetative
composition. These wetlands, together with
surrounding uplands, constitute the home range of
individual pairs. Usually, males become aggressive
toward other birds of the same species, defending
either wetlands within the home range or space
around their mates. These aggressive interactions
1
cause birds to distribute themselves throughout
the breeding habitat.
The need for dietary protein during the
prenesting and egg-laying periods causes ducks to
seek aquatic invertebrate foods, which may
compose 75 to 100% of the hen’s diet. Many species
maximize food acquisition during this period by
capitalizing on the seasonal peaks in aquatic food
abundance that differ among wetland types. For
example, shallow, temporary wetlands may exist
only a few weeks, but during that time they warm
quickly and develop invertebrate populations long
before permanent ponds. By moving among
wetlands and selecting those with the richest
invertebrate fauna, ducks are able to quickly
acquire the protein necessary for egg production.
Thus, small, shallow wetlands contribute as much
to ducks during the breeding period as large,
permanent cattail marshes. A diverse wetland
community is critical to this food acquisition
strategy.
Territorial aggression is often initiated when
males sight other birds of the same species. This
visual spacing limits the number of pairs that an
area can support. Habitats with many small ponds
on which ducks may isolate themselves, or those
with heavy vegetation, bays, or inlets where pairs
are visually separated, can reduce encounters
between birds and increase pair densities.
Wetlands most attractive to dabbling ducks contain
about a 50:50 ratio of open water to emergent
vegetation. Patches of emergent plants, sparse
enough to allow a duck to swim through, are more
attractive than large blocks of thick, unbroken
vegetation.
Nest Sites
Most diving ducks and some dabbling ducks
construct nests over water amid emergent
vegetation. In contrast, most dabbling duck nests
are made in dead vegetation remaining from the
previous growing season. Often, this residual
vegetation is found in grassland and shrub habitat
located up to a mile from water. Tall, dense grasses
or shrubs with low growth forms are usually
preferred by dabbling ducks. Islands also provide
attractive nesting habitat if adequate vegetative
cover is present. Hens explore many potential sites,
but select only one to construct a nest. Most ducks
lay a single egg each day until a clutch of 9 to 11
eggs is complete.
2 Fish and Wildlife Leaflet 13.2.7. • 1992
Incubation
As the clutch nears completion, hens begin an
incubation period that ranges from 23 to 30 days
for most species, with shorter periods typical of
species that lay smaller eggs. Duck nests are often
destroyed by mammalian, avian, or reptilian
predators. At present, throughout much of the
northern Great Plains, predators are abundant,
and duck nests are concentrated because nesting
cover is limited. Consequently, the percentage of
nests that hatch at least one egg (nest success) is
often less than 15%. In habitats where nests are
dispersed and predators are less common, much
higher (40 to 70%) success rates are typical. Most
ducks will renest if their initial clutch is destroyed
during laying or early in incubation and a
sufficient number and diversity of wetlands remain
available. In some species, hens that successfully
hatch a clutch often return to the vicinity of the
successful nest site in subsequent years, and
sometimes to the same nest bowl. During
incubation, hens leave the nest for a recess three to
five times per day. They continue to meet their
mates during these recesses until the male leaves
his territory and joins groups of other males in
preparation for molt. This usually occurs about 1 to
2 weeks into incubation.
Broods
Newly hatched ducklings leave the nest soon
after hatching, and may walk through uplands or
follow streams to brood-rearing wetlands up to a
mile away. Even after reaching a wetland, broods
may move among ponds. Ducklings of most species
feed almost entirely on aquatic invertebrates until
about a month old. Thereafter, ducklings of
dabbling duck species gradually increase their
consumption of seeds and other vegetation.
Because ducklings cannot thermoregulate until
they are about 2 weeks old, they are periodically
brooded by the hen. Predation and exposure can
cause high mortality among ducklings.
Contaminants can also cause mortality, either by
direct toxicity or, more often, by reducing the
abundance of essential invertebrate foods. In many
habitats, 20 to 50% of all duck broods are entirely
destroyed, and typically only about half of the
ducklings in the remaining broods survive. Habitat
use by broods differs among species, but is
generally related to the need for areas secure from
predators and severe weather. Diving duck broods
seek security in open water, where they dive to
escape predators. Dabbling duck broods usually
prefer dense emergent vegetation.
The Limiting Factor
Contemporary waterfowl management
generally uses three approaches for guiding
management actions. Actions initiated on an
international scale, such as in the North American
Waterfowl Management Plan, often originate from
broad policy directives such as the need to preserve
wetlands or increase nesting success. Other
initiatives are guided by computer simulations,
such as the Mallard Management Model, that
recommend actions based on knowledge of
waterfowl biology and factors that suppress
reproduction. However, similar guidelines are
generally unavailable for managing the scattered,
diverse duck breeding habitats of North America.
In such habitats, management actions are often
guided by the manager’s experience and intuition.
Predation, resource limitations, and
environmental conditions are factors that may
suppress waterfowl populations below their
biological potential. However, only one factor is
most limiting to populations at any time. Aldo
Leopold described the limiting factor as "the one
that has to be removed first, and usually the one to
which the application of a given amount of effort
will pay the highest returns, under conditions as
they stand." The effort required to remedy a
limiting factor may vary, but until it is removed,
activities directed at other, nonlimiting factors will
offer relatively little improvement in duck
production.
Although many contemporary ecologists view
the limiting factor concept as an oversimplification
of complex interrelationships, it is nonetheless a
useful starting point for considering factors that
suppress waterfowl recruitment. Sometimes, a
factor that limits duck production can result from
deficiencies independent of the breeding habitat,
for example, food shortages on wintering areas that
prevent the acquisition of fat reserves necessary for
successful breeding. Such limitations are usually
beyond the control of individual managers. Most
factors that are potentially limiting to duck
production, however, can be traced to four
important requirements of breeding habitat: the
ability to attract and retain spring migrants,
provide for the resource and social needs of
breeding pairs, secure adequate nesting habitat,
and provide suitable brood-rearing habitat.
Fish and Wildlife Leaflet 13.2.7. • 1992
Unfortunately, drought, localized agricultural
effects, and other dynamic events may cause
deficiencies in these requirements to vary annually.
Thus, management to correct long-term habitat
deficiencies should be based on average habitat
conditions. These average conditions should be
determined by evaluating premanagement
information collected during more than one
breeding season.
Because wetland communities are the basic
unit in which ducks live and acquire resources
during breeding, premanagement information
should be gathered independently for each discrete
community, not averaged across several isolated
wetland complexes. Although waterfowl
researchers are beginning to understand the
implications of habitat fragmentation for breeding
ducks, it is well established that the benefits of
small tracts of waterfowl habitat are often
swamped by the effects of habitat degradation on
adjacent lands. The protocol described here may
still be useful for identifying factors limiting duck
production, but management to overcome these
deficiencies on small tracts of land may be futile in
the face of overwhelming external forces.
Obtaining Premanagement
Information
Spring Migrants and Breeding Pairs
Information on the number of spring migrants
and resident breeding pairs can be obtained
through a series of ground counts beginning with
the first influx of spring migrants and continuing
through the early incubation period. Spring
migrant and pair counts, as well as brood counts,
should be conducted on a large block of contiguous
habitat that is representative of the management
area. Ideally, surveys should be conducted two or
three times per week, but in no case less than once
a week. Because females typically take incubation
recesses early and late in the day, nesting
chronology and indices to nest success are most
readily interpreted if observers restrict their
counts to the period between 1 hour after sunrise
to 1 hour before sunset. Observers should quietly
walk near wetlands but avoid flushing ducks. If
birds flush to nearby areas, observers should avoid
duplicate counts on these individuals. During the
time when spring migrants move through the
region, simply tally the numbers of individuals by
species and sex. When the number of ducks and the
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species composition stabilizes, one may assume
that many birds now in the area are beginning to
establish home ranges in preparation for breeding.
At this time, begin counting male-female pairs and
single males, tallying these males as "indicated
pairs." These single or "lone" males are usually
mates of females who are searching for nest sites,
laying eggs, or incubating. For each species, the
highest number of pairs plus indicated pairs
counted in any census represents the total
estimated pairs resident in the wetland community.
Nesting Habitat and Success
The quantity of available nesting habitat is
often easy to judge in relation to species
requirements. Most diving ducks construct nests
over water in robust emergent plants. Map the
distribution and vegetative composition of these
emergent beds, and note if such areas remain
inundated during the incubation period.
Cavity-nesting duck species use holes excavated by
woodpeckers or created by internal rot in old trees.
Note the number and distribution of potential nest
trees or actual nest sites and their distances from
the wetland. Dabbling ducks and some diving
ducks nest in grasses or shrubs adjacent to
wetlands. Map the area and distribution of these
habitats.
The quality of nesting habitat is difficult to
judge for overwater- and cavity-nesting species.
However, the height and density of upland sites
can be measured using a Robel pole or similar
device. Readings obtained at a standardized
viewing height and distance can then be compared
with minimum standards required by different
species. Whenever possible, managers should
determine the relative quality of potential nesting
habitat.
Duck nesting success is a more indirect index
of nesting habitat conditions because it is
dependent on the quality and quantity of habitat as
well as the density and composition of the local
predator community. In grassland habitats, large
numbers of nests can often be located using
cable-chain drags. In shrubland or wooded areas,
hand drags, dogs, or observations of hens returning
to nest sites may be necessary to locate nests.
When nests are found, note the size of the
completed clutch, candle the eggs to determine the
stage of incubation, then flag the site by placing a
marker at some set distance and direction away
from the nest. Excessive disturbance to the nest
site must be avoided. Later, revisit the site to
4 Fish and Wildlife Leaflet 13.2.7. • 1992
determine the fate of the nest. Nests that were
abandoned or destroyed by predators will contain
whole eggs and pieces of eggshell with membranes
firmly attached. Note the condition of the eggs and
look for tracks, scats, or other evidence that may
suggest the cause of nest failure. Successful nests
are typified by shell membranes that are easily
separated from shell fragments.
Brood-rearing Period
Begin duck brood surveys when broods of
early-nesting species first appear. Surveys should
be conducted in early morning (30 minutes before
to 1 hour after sunrise) and in late evening (2
hours before until 30 minutes after sunset). Counts
conducted at times other than early and late in the
day will census only a fraction of the broods
present and will be biased towards diving duck
species that use open water areas during
brood-rearing. Viewers should quietly observe
broods, from elevated vantage points if necessary,
and note the species, size of the brood (number of
ducklings), and age of the ducklings. Be aware that
duck broods may move among wetlands, and try to
avoid duplicate counts. If movements between
wetlands are uncommon and the number of broods
per wetland is low, it is often possible to distinguish
individual broods based on a combination of
species, size, and age. In such cases, note the
number of ducklings in a brood on subsequent
observations. If a brood is not observed on
subsequent surveys and the likelihood of secondary
movements to another rearing wetland is remote,
record the possibility that the entire brood
perished. To obtain data on duckling attrition,
individual broods should be observed every 3 to 5
days, particularly when ducklings are young and
mortality rates are highest. The most important
index to obtain during the brood-rearing period is
the number of young remaining in old (prefledging,
or class III) broods.
Identifying the Limiting Factor
Attracting and retaining spring migrants,
providing resources for breeding pairs, securing
adequate nesting habitat, and providing suitable
brood-rearing areas are all interdependent
activities, wherein each event is dependent on the
success of previous events. The following sections
provide a basis for identifying deficiencies in this
reproductive chain of events by interpreting the
Fig. 1. General management alternatives for addressing factors that limit duck recruitment. Readers should consult
technical publications for detailed information on specific alternatives.

Fish and Wildlife Leaflet 13.2.7. • 1992 5

premanagement data described above. Once a
limiting factor has been identified, general
management actions for correcting these
deficiencies can be considered (Fig. 1). Readers
should consult technical publications for
information on which management action is most
appropriate and how to implement an action.
Attracting and Holding Spring Migrants
and Breeding Pairs
Summarize data on the numbers of ducks
present in early spring, looking for evidence of a
sharp decline indicative of migrants departing the
area and resident pairs remaining behind. If large
numbers of migrants were present, but later
departed, and those migrants were species that
normally breed in the area, consider actions to
attract and hold spring migrants.
Examine the number of indicated breeding
pairs that remain after migrants leave the area,
then determine if the habitat is supporting
breeding pairs up to its potential. The key to
assessing this potential is knowing how many pairs
are attracted to good wetland communities in your
geographic area. Comparing pair densities on
nearby, high quality breeding habitat provides the
best basis for contrast. Historical data also can be
consulted. Lacking these data, managers should
consult state or federal agencies for area-specific
data. For example, curves depicting average
breeding pair densities as a function of wetland
size and type have been developed for the northern
Great Plains (e.g., Cowardin et al. 1988). Wetland
complexes that fail to attract adequate numbers of
breeding pairs can be managed to increase pair
numbers.
Enhancing Nesting Habitat and Nest
Success
Emergent vegetation suitable for overwater
nesters should be dense, have a height of at least 3
feet above water, and remain flooded during the
period of nesting. Suitable emergents should occur
in wide bands around the periphery of the wetland
or as large islands within the wetland basin. Most
cavity-nesting species select nest sites within 200
yards (183 m) of a wetland, although wood ducks
(Aix sponsa) will use cavities up to 1 mile (1.6 km)
from water. If suitable cavities are few or absent
within this area, artificial nesting structures can
help correct the deficiency. Ducks that nest in
upland sites require grasses, legumes, shrubs, or
6 Fish and Wildlife Leaflet 13.2.7. • 1992
combinations of the above plants within 1 mile of
wetlands. Suitable nesting areas should occur in
large (more than 40 acres or, 16 ha), unbroken
blocks of habitat.
Nesting cover should meet minimal Robel pole
indices for height and density (typically, dense at
heights of 18 inches—0.5 m—above the ground),
and should be secure from grazing and agricultural
manipulations until after the incubation period. If
density or height is insufficient, several
management actions can be used to enhance the
quality of nesting cover.
Data on the fate of marked nests should be
corrected for exposure, according to the Mayfield
correction technique, then average nest success
rates should be calculated for the management
area. Generally, nest success rates greater than
40% are acceptable in most habitats, whereas rates
lower than 15% are usually insufficient to maintain
a stable duck population. Lacking direct measures
of nest success, managers may obtain qualitative
indices of nest loss through "social indices" that
rely on the tendencies of many duck species to
renest if their initial nests are destroyed. The
simplest of these indices is an analysis of the
weekly ratios of indicated pairs (lone males) to
actual (male−female) pairs during the egg-laying
and incubation period for each species. Local
populations experiencing low rates of nest loss
often exhibit ratios that increase sharply in the
first few weeks, then gradually decline from a high
level (e.g., 0.2:1, 1.3:1, 3.4:1, 3.0:1, and 2.8:1).
Populations experiencing high nest loss may
exhibit an increase, followed by a sharp decrease,
then a subsequent increase in these ratios (e.g.,
0.2:1, 1.3:1, 3.4:1, 1.8:1, and 2.7:1), indicative of
unsuccessful hens rejoining their mates in
preparation for a second nesting attempt.
Additional evidence of nest destruction may be
derived by examining the hatching chronology of
duck broods for each species. This is accomplished
by back-dating broods to the date of hatch, using
information on duckling ages. A frequency
distribution of number of broods hatched within
5-day intervals typically depicts a peak of hatch
followed by a much smaller, well-defined, second
peak from renesting attempts (Fig. 2). Hatching
curves that exhibit pronounced renesting peaks or
are relatively flat suggest excessive rates of nest
loss.
If the quantity and quality of nesting cover are
adequate but nesting success is low, try to
determine the cause of nest failure. Predation is

Fig. 2. Hypothetical hatching curves for local duck
populations experiencing relatively high (top) and low
(middle and bottom) nesting success during early
incubation.
one common reason for nest failure in many
habitats, and may be indicated by evidence left at
the nest. However, do not discount the possibilities
of flooding, destruction from agricultural
operations, or exposure to weather. A wide array of
corrective actions are available to enhance nesting
success, depending on the cause of nest failure.
Improving Brood-rearing Habitat and
Duckling Survival
Duckling mortality is indicated either by loss of
complete broods or by brood attrition, wherein the
number of ducklings in a brood is reduced over
time. Mortality caused by exposure, starvation, or
death from pesticides or other contaminants often
results in the catastrophic loss of entire broods. In
contrast, mortality caused by predation may result
in a more gradual decrease in brood size. Generally,
an average of five ducklings per prefledging (class
III) brood is considered acceptable attrition.
Supplemental information, such as, from bait
Fish and Wildlife Leaflet 13.2.7. • 1992
stations to identify the presence of predators,
invertebrate sampling to gauge the abundance of
food, and water quality measures to detect
contaminants, may be needed to isolate the causes
of duckling mortality. Such supplemental data are
usually vital for selecting an appropriate
management strategy to enhance brood survival.
Rather than remain in undesirable habitat,
broods may move to other wetlands. The quality of
brood-rearing habitat may therefore be reflected by
the number of resident broods, compared with the
number of resident breeding pairs that were in the
area, after taking into account nest success rates
and renesting activity. If the estimated number of
broods occupying a wetland complex is far less than
the estimated number believed to have hatched,
management may be necessary to enhance the
quality of brood-rearing habitat. Often, the root
causes of low brood usage and poor brood survival
are the same, and a single management action may
be used to address both problems.
Other Considerations
Before initiating any management measure,
consider whether human disturbance or natural
forces have sufficiently altered the ecosystem to
warrant intervention. Do not use management
tools as "weapons" against a healthy landscape.
The waterfowl response to management of such
areas will be relatively slight when compared with
results of the same effort applied to dysfunctional
ecosystems. Unfortunately, however, some of the
most important waterfowl breeding habitats in
North America have been severely degraded. When
managing these habitats, overall objectives should
be consistent with the natural values of the
ecosystem. Not all wetlands are meant to be
breeding habitats. Migratory stopover and
wintering areas provide essential resources for
ducks, and managers should avoid modifying such
areas to create breeding habitat if doing so would
impair these other seasonal uses. Although
management actions can temporarily alter
waterfowl habitats for other than natural uses,
they do so only with high cost, intensive labor, and
possibly detrimental effects to the ecosystem.
Once a limiting factor has been identified and
an appropriate management response is devised,
managers should resist the temptation to
simultaneously initiate more than one action on a
single area. Imposing more than one management
treatment complicates evaluations of the
7
effectiveness of the actions, and often results in no
more success than a single treatment that is
selected with reasonable forethought.
Lastly, management actions should be
evaluated to determine whether the objectives of
the project were attained. The same techniques
and data analyses used when collecting
premanagement information should be employed
during this follow-up evaluation.
Suggested Reading
Bellrose, F. C. 1980. Ducks, geese, and swans of North
America. Stackpole Books, Harrisburg, Penn. 543 pp.
Cowardin, L. M., D. H. Johnson, T. L. Shaffer, and D. W.
Sparling. 1988. Applications of a simulation model to
decisions in mallard management. U.S. Fish and
Wildlife Service Technical Report 17. 28 pp.
Higgins, K. F., L. M. Kirsch, H. F. Duebbert, A. T. Klett,
J. T. Lokemoen, H. W. Miller, and A. D. Kruse. 1977.
Construction and operation of cable−chain drag for
nest searches. U.S. Fish and Wildlife Service
Wildlife Leaflet 512. 14 pp.
Johnson, D. H. 1979. Estimating nest success: the
Mayfield method and an alternative. Auk
96:651−661.
Kirby, R. E. 1988. American black duck breeding habitat
enhancement in the northeastern United States: a
review and synthesis. U.S. Fish and Wildlife Service
Biological Report 88(4). 50 pp.
Lokemoen, J. T. 1984. Examining economic efficiency of
management practices that enhance waterfowl
production. Transactions of the North American
Wildlife Natural Resources Conference 49:584−607.
Ringelman, J. K. 1991. Evaluating and managing
waterfowl habitat. Colorado Division of Wildlife
Division Report 16. 48 pp.
Robel, R. J., J. M. Briggs, A. D. Dayton, and L. C.
Hulbert. 1970. Relationships between visual
obstruction measurements and weight of grassland
vegetation. Journal of Range Management
23:295−297.
Weller, M. W. 1956. A simple field candler for waterfowl
eggs. Journal of Range Management 20:111−113.

Note: Use of trade names does not imply U.S. Government endorsement of commercial products.

UNITED STATES DEPARTMENT OF THE INTERIOR

FISH AND WILDLIFE SERVICE
Fish and Wildlife Leaflet 13
Washington, D.C. • 1992